KR100233191B1 - Chimeric gene for the transformation of plants - Google Patents

Chimeric gene for the transformation of plants Download PDF

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KR100233191B1
KR100233191B1 KR1019920003657A KR920003657A KR100233191B1 KR 100233191 B1 KR100233191 B1 KR 100233191B1 KR 1019920003657 A KR1019920003657 A KR 1019920003657A KR 920003657 A KR920003657 A KR 920003657A KR 100233191 B1 KR100233191 B1 KR 100233191B1
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르브룅 미쉘
르루 베르나르
싸일랑 알랭
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제닌 페트릭
롱-쁠랑 아그로
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    • C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
    • C12N15/8274—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for herbicide resistance
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    • C07K2319/01—Fusion polypeptide containing a localisation/targetting motif
    • C07K2319/08—Fusion polypeptide containing a localisation/targetting motif containing a chloroplast localisation signal

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Abstract

A new ADN transit peptide sequence comprises at least one coding sequence for the transit peptide from a plant gene encoding an enzyme located in plastids, the mature part of the N-terminal sequence of the plant gene coding for an enzyme located in plastids, and/or a sequence coding for a second transit peptide from a plant gene coding for an enzyme located in plastids. Independent claims are also included for the following: (1) a chimeric gene useful in plants to confer herbicidal resistance having an EPSPS target comprising a promoter region, a transit peptide region (as above), a sequence coding for resistance to glyphosate and a polyadenylated signal region not 3' translated; (2) a process for the construction of the above chimeric gene comprising isolating at least 2 regions of the transit peptide and at least part a sequence of the mature plant plastid genes to achieve at least one sequence coding for glyphosate resistance and a polyadenylated signal region therefore altogether providing plant resistance; (3) a vector for the transformation of the plant comprising the above chimeric gene; (4) a transformed plant cell containing the above chimeric gene; (5) a transformed plant with an improved resistance to herbicides having an EPSPS target obtained using the above cell; (6) a transformed plant with an improved resistance to herbicides having an EPSPS target stems from crossing the above plants; (7) a process for obtaining hybrid cell lines having glyphosate resistance using the above plants as parent plants; and (8) a process for treating the above plants comprising applying a herbicide having an EPSPS target.

Description

식물의 형질 전환용 키메라 유전자Plant chimera gene for transformation

본 발명은 신규의 전달 (transit)펩티드 DNA배열, 신규의 키메라 유전자, 및 일반적으로 제초제 특히 포스포노메틸글리신계의 제초제에 대한 식물의 내성을 증가시키기 위한 이들의 용도에 관한 것이다. 본 발명은 또한 이러한 유전자에 의해 형질 전환된 식물세포, 이 식물 세포로부터 재생된 형질전환 식물, 뿐만 아니라 이 형질 전환 식물을 이용한 이종교배에서 유래된 식물에 관한 것이다.The present invention relates to novel transit peptide DNA sequences, novel chimeric genes, and their use to increase plant resistance to herbicides, in particular herbicides, in particular phosphonomethylglycines. The present invention also relates to plant cells transformed with such genes, transgenic plants regenerated from these plant cells, as well as plants derived from crossbreeding using the transgenic plants.

글리포세이트, 술포세이트 또는 포사메틴은 포스포노메틸글리신계의 광역 스펙트럼계 제초제이다. 이들은 필수적으로 PEP(포스포에놀피루베이트)에 관련된 5-(에놀피루빌)쉬키메이트-3-포스페이트 합성효소(EC 2.5.1.19) 또는 EPSPS의 경합적인 저해제로서 작용한다. 상기 제초제가 식물에 적용된후, 제초제는 식물내부로 이동하여 급속성장부위, 특히 줄기 및 뿌리 정점에 축적되며, 민감성 식물의 퇴보 및 파괴를 일으킨다.Glyphosate, sulfosate or posamethine is a phosphonomethylglycine based broad spectrum herbicide. They essentially act as competing inhibitors of 5- (enolpyrubil) schikimate-3-phosphate synthetase (EC 2.5.1.19) or EPSPS, which are involved in PEP (phosphoenolpyruvate). After the herbicide is applied to the plant, the herbicide migrates inside the plant and accumulates at rapid growth sites, especially at the stem and root apex, causing degeneration and destruction of sensitive plants.

이들 제초제 제품의 주요 표적인 플래스티드(plastid) EPSPS는 방향족 아미노산 생합성 경로의 효소이며, 하나 이상의 핵 유전자에 의해 코팅되고 세포질내 전구체의 형태로 합성된 다음 플래스티드내로 들어와서 이곳에서 그의 천연 형태로 축적된다.The primary target of these herbicide products is the plastid EPSPS, an enzyme of the aromatic amino acid biosynthetic pathway, which is coated by one or more nuclear genes, synthesized in the form of precursors in the cytoplasm, and then into the plastid, in its natural form Accumulate.

글리포세이트 및 이 계통의 제품에 대한 식물의 내성은, EPSPS유전자의 생성물의 저해 특성에 대한 식물 또는 박테리아 유래의 돌연변이 또는 비돌연변이의 EPSPS유전자를 식물의 게놈에 안정하게 도입시킴으로써 얻어진다. 글리포세이트의 작용방식 및 사용되는 유전자의 생성물이 글리포세이트에 대해 갖는 내성도를 가정한다면, 상기 유전자의 번역생성물을 형질 발현시켜 이들을 플래스티드내에 실질적으로 축적시킬 수 있음을 이용할 수 있다.Plant resistance to glyphosate and products of this lineage is obtained by stably introducing into the genome of plants the mutant or non-mutant EPSPS genes derived from plants or bacteria to the inhibitory properties of the products of the EPSPS gene. Given the mode of action of glyphosate and the degree of resistance that the product of the gene used has to glyphosate, it can be utilized that the translation product of the gene can be expressed to substantially accumulate them in the plastid.

예를 들어 미합중국 특허 제4,535,060호에는, 플래스티드 구획내에 효소를 국재(localisation) 시킨후에, 경합적인 저해제(글리포세이트)에 대한 내성이 더욱 큰 효소를 형성하는 적어도 하나의 돌연변이 보유 EPSPS를 코딩하는 유전자를 식물게놈내에 도입함으로써, 상기 언급된 유형의 제초제, 특히 N-(포스포노메틸)글리신 또는 글리포세이트에 대한 내성을 식물에 부여하는 것이 공지되어 있다. 그러나, 이 기술은 농경 조건하에서 식물을 이용함에 있어서 더욱 큰 신뢰성을 획득하기 위해서는 개량이 필요하다.For example, US Pat. No. 4,535,060 describes at least one mutant bearing EPSPS that forms an enzyme with greater resistance to competitive inhibitors (glyphosate) after localization of the enzyme in the plastid compartment. By introducing genes into plant genomes, it is known to confer plants to resistance to the above-mentioned herbicides, in particular N- (phosphonomethyl) glycine or glyphosate. However, this technique requires improvement to obtain greater reliability in using plants under agricultural conditions.

본 명세서에서, “식물”이란 광합성이 가능한 분화된 다세포 유기체를 의미하며, “식물세포”란 유합조직(callus)과 같은 비분화 조직 또는 배 또는 식물 마디, 식물 또는 종자와 같은 분화 조직을 형성할 수 있는 식물에서 유래된 세포를 의미한다.As used herein, "plant" means a differentiated multicellular organism capable of photosynthesis, and "plant cells" may form undifferentiated tissues such as callus or differentiated tissues such as pears or plant nodes, plants or seeds. It means a cell derived from a plant that can.

본 발명의 목적은, 제초제에 대한 내성이 있는 유전자를 함유하는 신규의 키메라유전자에 의해 형질전환된 세포를 재생시킴으로써, 일반적으로 제초제, 특히 포스포노메틸글리신계의 제초제에 대한 내성이 증가된 형질 전환 식물을 생성하는데 있다. 본 발명은 또한 신규의 키메라유전자, 이를 함유하는 신규의 전달 펩티드, 뿐만아니라 식물내에 돌연변이 효소가 그의 성숙형으로 축적됨으로써 더욱 내성이 증가된 식물에 관한 것이다.An object of the present invention is to transform cells that have been transformed by novel chimeric genes containing genes that are resistant to herbicides, thereby generally increasing the resistance to herbicides, in particular phosphonomethylglycine-based herbicides. To produce plants. The present invention also relates to novel chimeric genes, novel delivery peptides containing them, as well as plants that are further tolerated by the accumulation of mutant enzymes in the plant in their mature form.

더욱 특별하게는, 본 발명의 목적은 전사방향으로, 프로모터 영역, 전달 펩티드 부위, 글리포세이트 내성 효소를 코딩하는 유전자 배열 및 3′의 비번역 폴리아데닐화 시그널 영역을 함유함을 특징으로 하는, 표적이 EPSPS인 제초제에 대해 증가된 내성을 식물에 부여하는 키메라유전자이며, 상기 전달 펩티드 부위는 그의 전사 방향으로 플래스티드-위치 효소를 코딩하는 식물 유전자의 전달 펩티드, 플래스티드-위치 효소를 코딩하는 식물 유전자의 N-말단 성숙부분의 부분적 배열, 및 플래스티드-위치 효소를 코딩하는 식물 유전자의 두번째 전달 펩티드를 함유하고 있다.More particularly, the object of the present invention is characterized by containing in the transcription direction a promoter region, a delivery peptide region, a gene sequence encoding a glyphosate resistant enzyme and a 3 ′ untranslated polyadenylation signal region, A chimeric gene conferring increased resistance to a herbicide whose target is EPSPS, wherein the delivery peptide site encodes a delivery peptide of the plant gene, the plastid-position enzyme, that encodes the plastid-position enzyme in its transcription direction A partial arrangement of the N-terminal mature portion of the plant gene, and a second delivery peptide of the plant gene encoding the flashed-position enzyme.

본 발명은 또한 상기 정의된 전달 펩티드 영역의 DNA배열에 관한 것이다.The invention also relates to the DNA sequence of the delivery peptide region as defined above.

전달 펩티드 부위에서 사용될 수 있는 전달 펩티드는 그 자체로 공지될 수 있으며, 예를 들어 옥수수, 해바라기, 완두콩, 담배 등에서 유래된 식물 기원의 것일 수도 있다. 첫번째 및 두번째 전달 펩티드는 동일하거나 유사하거나 상이할 수도 있다. 이들은 또한 각각 하나이상의 전달 펩티드 단위를 함유할 수도 있다. 리불로오스 1,5-디포스페이트 카르복실라아제 옥시게나아제(RuBisCO)유전자의 SSU에서 유래된 배열이 바람직하게 사용된다.The delivery peptides that can be used at the delivery peptide sites can be known per se and can be of plant origin derived from, for example, corn, sunflower, peas, tobacco, and the like. The first and second delivery peptides may be the same, similar or different. They may also each contain one or more delivery peptide units. An arrangement derived from the SSU of the ribulose 1,5-diphosphate carboxylase oxygenase (RuBisCO) gene is preferably used.

N-말단 성숙부분의 부분적 배열은 옥수수, 해바라기 또는 완두콩 유전자등과 같은 플래스티드-위치 효소를 코딩하는 식물유전자에서 유래되며, 기원의 식물 종은 서로 동일하거나 유사하거나 상이할 수 있고 이로 부터 첫번째 및 두번째 전달 펩티드가 각각 유래된다. 또한, 성숙 부분의 부분적 배열은 다양한 수, 일반적으로 10∼40, 바람직하게는 18∼33개의 아미노산을 함유할 수도 있다. 리불로오스 1,5-디포스페이트 카르복실라아제 옥시게나아제(RuBisCO) 유전자의 SSU에서 유래된 배열이 바람직하게 사용된다.The partial arrangement of the N-terminal mature part is derived from plant genes encoding plastid-position enzymes such as the corn, sunflower or pea genes, and the plant species of origin may be identical, similar or different from each other and the first and Each second delivery peptide is derived. The partial arrangement of the mature moiety may also contain various numbers, generally 10-40, preferably 18-33 amino acids. An arrangement derived from SSU of the ribulose 1,5-diphosphate carboxylase oxygenase (RuBisCO) gene is preferably used.

전체 전달 부위의 구성은 공지된 방법, 특히 융합 또는 임의의 기타 적절한 수단에 따라 수행될 수도 있다. 이 특징적 영역의 역할은 성숙한 천연 단백질을 최대 효율로 방출시킬 수 있는 것이다.The construction of the entire delivery site may be carried out according to known methods, in particular fusion or any other suitable means. The role of this characteristic region is to be able to release mature natural protein with maximum efficiency.

본 발명에 따른 키메라 유전자에서 사용될 수도 있는 제초제 내성을 위한 코드배열은 글리포세이트 내성을 갖는 돌연변이 EPSPS를 코딩한다. 특히 EPSPS유전자의 돌연변이에 의해 수득되는 이 배열은 박테리아 기원 예를 들어 살모넬라 티피뮤리움(Salmonella typhymurium) 유래(이하 “AroA 유전자” 라고 함) 또는 식물 기원, 예를 들어 페튜니아 또는 토마토 유래일 수도 있다. 이 서열은 하나 이상의 돌연변이, 예를 들어 Pro 101의 Ser으로의 돌연변이 또는 대안적으로 Gly 96의 Ala으로의 돌연변이를 함유할 수도 있다.The coding sequence for herbicide tolerance, which may be used in the chimeric gene according to the invention, encodes a mutant EPSPS with glyphosate resistance. This arrangement, in particular obtained by mutation of the EPSPS gene, may be of bacterial origin, for example Salmonella typhymurium (hereinafter referred to as “AroA gene”) or of a plant origin, for example petunia or tomato. This sequence may contain one or more mutations, eg, a mutation of Pro 101 to Ser or alternatively a mutation of Gly 96 to Ala.

본 발명에 따른 키메라 유전자의 프로모터 부위는 유리하게는 식물에서 자연적으로 형질 발현되는 유전자의 단편상의 적어도 하나의 프로모터, 다시 말해서 콜리플라워 모자이크 비루스의 35S RNA(CaMV35S)의 프로모터와 같은 비루스 유래의 프로모터, 또는 옥수수 또는 해바라기와 같은 곡류의 리불로오스 1,5-디포스페이트 카르복실라아제(RuBisCO) 유전자의 작은 서브 유니트와 같은 식물 기원의 프로모터로 구성될 수도 있다.The promoter region of the chimeric gene according to the invention advantageously comprises at least one promoter on a fragment of a gene that is naturally expressed in a plant, ie a promoter derived from a virus, such as a promoter of 35S RNA (CaMV35S) of cauliflower mosaic virus, Or promoters of plant origin, such as small subunits of the ribulose 1,5-diphosphate carboxylase (RuBisCO) gene of cereals such as corn or sunflower.

본 발명에 따른 키메라 유전자의 3′의 비번역 폴리아데닐화 시그날 영역은 예를 들어 노팔린 합성 효소 유전자와 같은 박테리아 기원, 또는 옥수수 또는 해바라기 RuBisCO의 작은 서브유니트와 같은 식물기원 유래일 수도 있다.The 3 ′ untranslated polyadenylation signal region of the chimeric gene according to the invention may be of bacterial origin, eg a plant origin, such as a small subunit of corn or sunflower RuBisCO, for example.

본 발명에 따른 키메라 유전자는, 상기 필수 부위외에 추가로, 프로모터 부위와 코드 배열 사이에 박테리아, 비루스 또는 식물 유래의 것일 수도 있는 비번역 중간 부위(링커)를 함유할 수도 있다.In addition to the essential site, the chimeric gene according to the present invention may further contain an untranslated intermediate site (linker), which may be of bacterial, viral or plant origin, between the promoter site and the code sequence.

[실시예 1]Example 1

[키메라 유전자의 구성][Configuration of Chimeric Genes]

본 발명에 따른 키메라 유전자의 구성은 하기 요소를 사용하여 수행된다:The construction of chimeric genes according to the invention is carried out using the following elements:

1) “이중 CaMV”프로모터(다시 말해서, 중복되는 부분)1) “Dual CaMV” promoter (ie, overlapping part)

CaMV 35S프로모터를 오델(Odell)등의 방법(1985)에 의해 단리한다. 전사 개시 부위의 상류쪽에 약 850bp을 함유하는 클론 pJO 5-2을 EcoRI- HindIII로 절단하고, 단리된 단편의 끝을 클레노우 폴리머라아제(Klenow polymerase)를 사용하여 블런트화 하고, 단편을 벡터 pUC 19(야니시-페론(Yannish-Perron)등, 1985)의 HincII 부위에 삽입한다.The CaMV 35S promoter is isolated by the method of Odell et al. (1985). Cloning pJO 5-2 containing about 850 bp upstream of the transcription initiation site was digested with EcoRI- HindIII, the ends of the isolated fragments blunted using Klenow polymerase, and the fragments were vector pUC. It is inserted into the HincII site of 19 (Yannish-Perron et al., 1985).

이 프로모터를 ClaI으로 소화하고, 말단을 클레노우 폴리머라제를 사용하여 채운다음 HindIII로 재소화한다. 동일한 초기 프로모터로부터 단리된 HindIII-EcoRV단편을 이들 두 부위 사이에 도입한다. 이렇게 수득된 펩티드는 CaMV35S 프로모터의 조절 요소의 이중 증폭 영역 상류를 갖는다. 이것을 HindIII-EcoRI단편 형태로 프랑스공화국 특허 출원 88/04130에 기재되어 있고, HindIII 및 EcoRI으로 절단된 벡터 pRPA-BL 150A 알파 2내로 도입한다.This promoter is digested with ClaI, the ends are filled with Clenow polymerase and then digested with HindIII. HindIII-EcoRV fragments isolated from the same initial promoter are introduced between these two sites. The peptide thus obtained has a double amplification region upstream of the regulatory element of the CaMV35S promoter. This is described in French patent application 88/04130 in the form of a HindIII-EcoRI fragment and introduced into the vector pRPA-BL 150A alpha 2 cleaved with HindIII and EcoRI.

2) 전이영역(Transfer region) : 두개의 전달 펩티드 뿐만이 아니라 사용된 성숙 단백질 요소는 그의 유전자가 리브룬(Lebrun)등 (1987)에 의해 기술된 옥수수 RuBisCO의 작은 서브유니즈의 클론화 cDNA및 왁스만(Waksman)등 (1987)에 의해 단리된 해바라기 RuBisCO의 작은 서브유니트의 클론화된 cDNA로 부터 얻는다. 보다 구체적으로, 전달 영역은 최적화 전달 펩티드로 명명되며 번역 방향으로 하기를 포함한다:2) Transfer region: The two transfer peptides as well as the mature protein elements used were cloned cDNAs and waxes of a small subunit of maize RuBisCO whose genes were described by Lebrun et al. (1987). From cloned cDNA of a small subunit of sunflower RuBisCO isolated by Waksman et al. (1987). More specifically, the delivery region is termed an optimized delivery peptide and includes the following in the direction of translation:

- 해바라기 RuBisCO의 작은 서브유니트의 전달 펩티드,-Delivery peptide of small subunits of sunflower RuBisCO,

- 옥수수 RuBisCO의 작은 서브유니트의 성숙 부분의 22개 아미노산의 N-말단 배열,N-terminal arrangement of 22 amino acids of the mature part of a small subunit of corn RuBisCO,

- 옥수수 RuBisCO 의 작은 서브유니트의 전달 펩티드,-Corn RuBisCO, a delivery peptide of small subunits,

이 최적화 전달펩티드를 사용한 구성물은 pRPA-BL로 명명한다.The construct using this optimized delivery peptide is named pRPA-BL.

10 내지 40개의, 바람직하게는 18 내지 33개의 아미노산 배열을 각각 함유하는 다른 유사한 배열이 사용될 수 있다.Other similar arrangements may be used, each containing 10 to 40, preferably 18 to 33, amino acid sequences.

대조용 요소를 제공하기 위하여, 선행기술에 따라서 첫번째 전달펩티드 및 두번째 성숙 배열을 사용하나 두번째 전달 펩티드를 사용하지 않고, 또 다른 구성을 실행한다(pRPA-BL 294).To provide a control element, another configuration is performed according to the prior art using the first delivery peptide and the second maturation sequence but without the second delivery peptide (pRPA-BL 294).

3) 구조유전자 : 스탈커(Stalker)등 (1985)에 의해 단리된 살모넬라 티피뮤리움의 EPSPS의 위치(Pro 101가 Ser로 됨)에서 돌연변이 유전자를 얻는다. pMG 34-2 클론(Calgene 공급)을 XbaI으로 선상화한 다음 비그나 라디아타(Vigna radiata) 뉴클레아제로 처리한다. SmaI으로 재절단한 후에, 두개의 블런트 말단을 연결한다. 수득된 클론은 정지 코돈의 하류의 17-bp SalI부위 뿐만아니라 개시코돈 ATG내의 NcoI부위를 갖는다. 이 클론을 pRPA-BL 104로 명명한다.3) Structural gene: The mutant gene is obtained at the position of EPSPS (Pro 101 becomes Ser) of Salmonella typhimurium isolated by Stalker et al. (1985). pMG 34-2 clone (Calgene supply) is linearized with XbaI and then treated with Vigna radiata nuclease. After recutting with SmaI, connect the two blunt ends. The clone obtained had the NcoI site in the start codon ATG as well as the 17-bp SalI site downstream of the stop codon. This clone is named pRPA-BL 104.

4) 폴리아데닐화 시그널 영역 : 이 단편은 pTi 37의 노팔린 합성 효소 유전자(베반(Brvan)등, 1983)로 부터 얻는다. 이 부위는 BamHI 및 EcoRI부위를 각각 5′ 및 3′ 말단에 도입하기 위하여 클레노우 폴리머라제로 처리하고 Ml3 mp 18의 SmaI부위에서 클론화한 260-bp MboI 단편(프랄리(Fraley)등, 1983; 특허 출원 PCT 84/02913)내에 함유되어 있다.4) Polyadenylation Signal Region: This fragment is obtained from the nopaline synthase gene of pTi 37 (Brvan et al., 1983). This site was a 260-bp MboI fragment (Fraley et al., 1983 treated with Clenow polymerase and cloned from the SmaI site of Ml3 mp 18 to introduce BamHI and EcoRI sites at the 5 ′ and 3 ′ ends, respectively. The patent application PCT 84/02913.

BamHI으로 절단하고 비그나 라디아타 뉴클레아제로 처리한 후에 EcoRI으로 절단하고 클레노우 폴리머라제로 처리하여, 생성 단편을 벡터 p-BL 20(참고. 프랑스공화국 특허출원 88/04130)내에 도입하고, XbaI 및 BamHI으로 절단하고 클레노우 폴리머라제로 처리한다. SalI 및 SstI으로 재절단한 후에, SalI부위면상에 3′nos 배열 및 SstI부위의 T-DNA면상에 오른쪽말단을 함유하는 약 0.4kbp의 단편이 수득된다.Cleavage with BamHI, treatment with Vigna radiata nuclease, cleavage with EcoRI, treatment with cleno polymerase, introduce the resulting fragments into vector p-BL 20 (see French Patent Application 88/04130) and XbaI And BamHI and treated with Klenow polymerase. After recutting to SalI and SstI, a fragment of about 0.4kbp was obtained containing a 3'nos arrangement on the SalI site and the right end on the T-DNA plane of the SstI site.

이들 모든 요소의 구성을 하기 방법으로 실행한다 :Perform the configuration of all these elements in the following way:

“옥수수 RuBisCO 의 SSU의 전달 펩티드/AroA 유전자” 융합 :"Transfer Peptide / AroA Gene of SSU in Corn RuBisCO" Fusion:

옥수수 RuBisCO 유전자의 SSU의 전달 펩티드는 전달 펩티드의 분해 부위에 상응하는 SpI 및 번역 개시 코돈을 걸치는 NcoI부위를 갖는, 레브룬 등(1987)에 의해 기술된 옥수수 RuBisCO유전자의 SSU유전자에 상응하는 cDNA로 부터 수득한 192-bp EcoRI-SphI 단편으로 부터 얻는다.The delivery peptide of the SSU of the maize RuBisCO gene is the cDNA corresponding to the SSU gene of the maize RuBisCO gene described by Revrun et al. (1987), with SpI corresponding to the cleavage site of the delivery peptide and the NcoI site spanning the translation initiation codon. From 192-bp EcoRI-SphI fragment obtained from

번역 융합물은 옥수수 전달 펩티드 및 세균 EPSPS 유전자 사이에서 SphI 말단을 박테리오파아지 T4 폴리머라제로 처리하고, EcoRI으로 재절단된, pRPA-BL 104로 부터의 AroA유전자의 클레노우 폴리머라제-처리 NcoI 말단과 연결함으로써 수득된다.The translational fusions were treated with the bacteriophage T4 polymerase at the SphI terminus between the corn transit peptide and the bacterial EPSPS gene and with the Klenow polymerase-treated NcoI terminus of the AroA gene from pRPA-BL 104, recut with EcoRI. It is obtained by connecting.

“옥수수 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 성숙부분의 22개 아미노산의 배열/AroA 유전자” 융합 :"Transfer Peptides of SSU in Corn RuBisCO / Arrangement of 22 Amino Acids / AroA Gene of Mature Part of Corn RuBisCO" Fusion:

유사하게, 옥수수 RuBisCO유전자의 SSU의 cDNA의 228-bp EcoRI-HindII 단편을 pRPA-BL 104 로 부터의 AroA 유전자의 클레노우 폴리머라제-처리 NcoI 말단과 연결하고 EcoRI으로 재절단한다. 번역 융합물은 옥수수 RuBisCO의 SSU의 전달 펩티드, 옥수수 RuBisCO의 SSU의 성숙부분의 22개 아미노산 및 세균 EPSPS유전자사이에서 수득된다.Similarly, the 228-bp EcoRI-HindII fragment of the cDNA of SSU of maize RuBisCO gene is linked to the Klenow polymerase-treated NcoI end of the AroA gene from pRPA-BL 104 and re-cut into EcoRI. Translation fusions are obtained between the delivery peptides of the SSU of maize RuBisCO, the 22 amino acids of the mature portion of the SSU of maize RuBisCO and the bacterial EPSPS gene.

해바라기 RuBisCO의 SSU의 전달 펩티드 :Sunflower RuBisCO SSU Delivery Peptides:

단편을 왁스만(Waksman) 및 프레이지네트(Freyssinet)(1987)에 의해 단리된 cDNA로 부터 얻는다. SphI부위가 졸러(Zoller)및 스미쓰(Smith)(1984)의 방법에 따라서 전달 펩티드의 분해 분위에서 만들어진다. 이렇게 수득된 해바라기 RuBisCO의 SSU의 전달 펩티드는 171-bp EcoRI-SphI 단편이다.Fragments are obtained from cDNAs isolated by Waksman and Freysinet (1987). SphI sites are made at the cleavage site of the delivery peptide according to the method of Zoler and Smith (1984). The delivery peptide of the SSU of sunflower RuBisCO thus obtained is 171-bp EcoRI-SphI fragment.

“해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 SSU의 성숙부분의 22개 아미노산의 배열/AroA 유전자” 융합 :“Transfer Peptides of Sunflower RuBisCO SSU / Amino Acid Arrangement / AroA Gene of Mature Part of SSU of Corn RuBisCO” Fusion:

옥수수 RuBisCO의 SSU의 전달 펩티드/옥수수 유전자 융합물의 성숙 부분의 옥수수 RuBisCO의 SSU의 22개 아미노산의 배열을 함유하는 구성물을 해바라기 RuBisCO의 SSU의 전달 펩티드에 상응하는 171-bp EcoRI-SphI으로 절단한다. 생성 구성물은 EcoRI-SphI 단편의 치환을 보여주며 이것은 번역 융합물 “해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 SSU의 성숙 부분의 22개 아미노산의 배열/AroA 유전자”이다.The construct containing the 22 amino acid sequence of SSU of maize RuBisCO of the mature portion of the delivery peptide / corn gene fusion of SSU of maize RuBisCO is cleaved with 171-bp EcoRI-SphI corresponding to the delivery peptide of SSU of maize RuBisCO. The resulting construct shows the substitution of the EcoRI-SphI fragment, which is a translational fusion “A delivery peptide of the SSU of sunflower RuBisCO / Sequence of 22 amino acids / AroA gene of the mature part of the SSU of corn RuBisCO”.

EcoRI-SalI 단편을 3′ nos 배열 및 T-DNA 의 오른쪽 말단을 함유하는 SalI-SstI단편과 연결한다. “해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 SSU의 성숙 부분의 22개 아미노산의 배열/AroA유전자 /3′ nos/T-DNA오른쪽 말단”으로 구성된 생성 EcoRI-SstI 단편을 이중 CaMV 프로모터를 함유하는 플라스미드 150A 알파 2의 T-DNA의 오른쪽 말단을 함유하는 EcoRI-SstI 단편으로 치환한다. 벡터 150A알파 2내의 전사 융합물 “이중 CaMV/해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 SSU의 성숙 부분의 22개 아미노산의 배열/AroA 유전자/3′ nos”를 pRPA-BL 294로 명명한다.The EcoRI-SalI fragment is linked to the SalI-SstI fragment containing the 3 ′ nos sequence and the right end of the T-DNA. The resulting EcoRI-SstI fragment consisting of “an array of 22 amino acids / AroA gene / 3 ′ nos / T-DNA right end” of the delivery peptide of the sunflower RuBisCO SSU / corn mature part of the RuBisCO SSU / AroAgene / 3′nos / T-DNA containing a double CaMV promoter Substitute the EcoRI-SstI fragment containing the right end of the T-DNA of plasmid 150A alpha 2. The transcriptional fusion in vector 150A alpha 2 “Double CaMV / Delivery Peptide of SSU of Sunflower RuBisCO / Sequence of 22 amino acids / AroA gene / 3 ′ nos of mature portion of SSU of Corn RuBisCO” is named pRPA-BL 294.

“해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 RuBisCO의 SSU의 22개 아미노산의 배열/옥수수 RuBisCO의 SSU의 전달 펩티드/AroA 유전자” 융합 :"Sweeping Peptides of Sunflower RuBisCO / Su Peptides of Corn RuBisCO SSU / Supplementing Peptides / AroA Genes of Corn RuBisCO SSU" Fusion:

상기 구성물을 NcoI-HindIII로 절단하여, AroA 유전자를 분리한다. 다음에, “옥수수 RuBisCO의 SSU의 전달펩티드/ AroA 유전자” 융합물을 함유하는 1.5kbp NcoI-HindIII 단편과 연결한다. 생성 구성물은 NcoI-HindIII 단편의 치환을 보여주며, 이것은 전사 융합물 “해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 유전자의 성숙 부분의 RuBisCO의 SSU의 22개 아미노산의 배열/옥수수 RuBisCO의 SSU의 전달펩티드/ AroA 유전자” 이다.The construct is cleaved with NcoI-HindIII to isolate the AroA gene. Next, it is linked with a 1.5kbp NcoI-HindIII fragment containing the "transpeptide / AroA gene of SSU of Corn RuBisCO" fusion. The resulting construct shows the substitution of the NcoI-HindIII fragment, which indicates the transcriptional fusion "peptide of the SSU of the RuBisCO / array of 22 amino acids of the RuBisCO SSU of the mature part of the sunflower gene RuBisCO / of the SSU of the corn RuBisCO / AroA gene ”.

EcoRI-SalI 단편을 3′ nos 배열 및 T-DNA의 우측 말단을 함유하는 SalI-SstI 단편과 연결한다. “해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 유전자의 성숙 부분의 RuBisCO의 SSU의 22개 아미노산의 배열/옥수수 RuBisCO의 SSU의 전달펩티드/ AroA유전자/3′ nos/T-DNA우측말단”을 포함하는 생성 EcoRI-Sst 단편이 이중 CaMV프로모터를 함유하는 플라스미드 150A 알파 2의 T-DNA의 우측 말단을 함유하는 EcoRI-SstI으로 치환된다. 벡터 150A 알파 2내의 전사 융합물 “이중 CaMV/해바라기 RuBisCO의 SSU의 전달 펩티드/옥수수 유전자의 성숙 부분의 RuBisCO의 SSU의 22개 아미노산의 배열/옥수수 RuBisCO의 SSU의 전달펩티드/ AroA 유전자/3′ nos” 를 pRPA-BL 410으로 명명한다.The EcoRI-SalI fragment is linked with the SalI-SstI fragment containing the 3 ′ nos configuration and the right end of the T-DNA. Production comprising "arrangement of 22 amino acids of RuBisCO SSU / delivery peptide of corn RuBisCO SSU / AroA gene / 3 'nos / T-DNA right end" of the mature peptides of the sunflower RuBisCO SSU delivery peptide / corn gene The EcoRI-Sst fragment is substituted with EcoRI-SstI containing the right end of the T-DNA of plasmid 150A alpha 2 containing the double CaMV promoter. Transcription Fusion in Vector 150A Alpha 2 “Double CaMV / Delivery Peptides of Sunflower SSU of Peptides / Massum RuBisCO / Arrangement of 22 amino acids of RuBisCO SSU of Peptides / Massage of Maize / BuR RuBisCO SSU Peptides / AroA Genes / 3 ′ nos Is named pRPA-BL 410.

[실시예 2]Example 2

[형질전환된 식물의 내성]Tolerance of Transgenic Plants

1. 형질전환 :1. Transformation:

벡터를 코스미드 pTVK 291(코마리(Komari)등, 1986)을 운반하는 노논코제닉 아그로박테리움(nononcogenic agrobacterium)균주 EHA 101(후드(Hood)등, 1987) 내로 도입한다. 형질전환 방법은 호쉬(Horsh) 등의 방법(1985)을 기초로 한다.The vector is introduced into nononcogenic agrobacterium strain EHA 101 (Hood et al., 1987) carrying cosmid pTVK 291 (Komari et al., 1986). The transformation method is based on the method of Horsh et al. (1985).

2. 재생 :2. Play:

담배 PBD 6(공급 SEITA, 프랑스)의 재생을 잎 외식체를 사용하여 30g/l의 수크로오즈 및 200g/ml의 카나마이신을 함유하는 무라쉬지(Murashige) 및 스쿠그(Skoog)(MS)염기성 배지상에서 실행한다. 잎 외식체를 온실 또는 시험관내-성장 식물로 부터 제거하고 잎 원반조직 방법(foliar disc method)(Science 1985, Vol. 227, p1229-1231)에 따라서 일련의 삼단계로 형질전환시킨다: 첫번째는 0.05mg의 나프틸아세트산(ANA)및 2mg/l의 벤질아미노퓨린(BAP)을 함유하는 30g/l의 수크로오즈로 보강된 MS 배지상에서 15일간 싹을 유도하는 것이다. 이 단계에서 형성된 싹은 30g/l의 수크로오즈로 보강은 되었으나 호르몬은 함유하지 않는 MS배지상에서 배양함으로써 성장된다. 성장된 싹을 제거하고 이것을 1/2양의 염, 비타민 및 당을 함유하나 호르몬을 함유하지 않는 1/2-희석된 MS 이식 배지상에서 배양한다. 약 15일후에, 뿌리가 깊게 자란 싹을 토양에 심는다.Regeneration of tobacco PBD 6 (feed SEITA, France) using leaf explants Murashige and Skoog (MS) basic medium containing 30 g / l sucrose and 200 g / ml kanamycin Run on Leaf explants are removed from the greenhouse or in vitro-growing plants and transformed in a series of three steps according to the leaf disc method (Science 1985, Vol. 227, p1229-1231): first 0.05 Shoots are induced for 15 days on MS medium supplemented with 30 g / l sucrose containing mg of naphthyl acetic acid (ANA) and 2 mg / l benzylaminopurine (BAP). The shoots formed at this stage are grown by culturing on MS medium supplemented with 30 g / l sucrose but not containing hormones. The grown shoots are removed and cultured on 1 / 2-diluted MS transplant medium containing 1/2 amount of salt, vitamins and sugars but no hormones. After about 15 days, deep-rooted shoots are planted in the soil.

3. 글리포세이트 내성의 측정 :3. Measurement of glyphosate resistance:

a) 시험관내 : 35ppm의 글리포세이트 및 200마이크로그램/ml의 카나마이신을 함유하는 30g/l의 수크로오즈, 0.05mg/l의 나프탈렌 아세트산 및 2mg/l의 BAP로 보강된 MS 배지상에서 30일간 성장한 후의, 0.5cm직경의 100개의 잎이 원반상 조직으로 추정한 유합조직의 중량을 측정함으로써 내성을 측정한다. 이들 조건하에서, 본 발명에 따른 pRPA Bl 410의 키메라 유전자에 의해 변성된 담배 식물에 있어서, 유합조직의 중량은 34g의 반면에 두번째 전달펩티드가 없는 키메라유전자에 의해 변성된 식물에 대한 유합조직의 중량은 단지12g이다.a) in vitro: 30 days in MS medium supplemented with 30 g / l sucrose containing 35 ppm glyphosate and 200 micrograms / ml kanamycin, 0.05 mg / l naphthalene acetic acid and 2 mg / l BAP After growth, 100 leaves of 0.5 cm diameter were measured for resistance by measuring the weight of the callus tissue estimated to be discoid tissue. Under these conditions, in tobacco plants denatured by the chimeric gene of pRPA Bl 410 according to the present invention, the weight of the callus was 34 g while the weight of callus to the plant denatured by the chimeric gene without the second delivery peptide. Is only 12g.

b) 생체내 : pRPA-BL 294 및 pRPA-BL 410을 각각 사용하여 형질전환된 담배를 재생하여 얻은 30개의 식물을 온실로 옮기고 5-잎 단계에서 0.6kg/ha의 글리포세이트(Round up) 에 상응하는 투여량으로 수성 현탁액을 분무한다. 21일 후에 형질전환 되지 않은 대조용 식물에 대한 식물의 표현형 시험을 실행한다. 이들 조건하에서, pRPA-BL 410을 사용하여 형질전환된 식물은 무시할만한 식물 독성을 갖는 반면에 대조용 식물은 완전히 파괴되었다. 또한, 두번째 전달 펩티드가 없다는 것이 상기 펩티드와 다른 키메라 유전자를 사용하여 형질전환된 식물은 30% 이하로 파괴되는 식물 독성을 갖는다.b) In vivo: 30 plants obtained by regenerating transformed tobacco using pRPA-BL 294 and pRPA-BL 410 respectively were transferred to a greenhouse and rounded up to 0.6 kg / ha of glyphosate at the 5-leaf stage. Spray the aqueous suspension at a dose corresponding to that. After 21 days a phenotypic test of the plants is performed on the control plants that were not transformed. Under these conditions, plants transformed with pRPA-BL 410 had negligible phytotoxicity while control plants were completely destroyed. In addition, the absence of a second delivery peptide has plant toxicity in which plants transformed with the peptide and other chimeric genes are destroyed up to 30%.

이들 결과는 본 발명에 따른 글리포세이트 내성을 코딩하는 키메라 유전자를 사용함으로써 명백히 개선되었음을 보여준다.These results show a clear improvement by using a chimeric gene encoding glyphosate resistance according to the present invention.

본 발명에 따라서 형질전환된 균주는 글리포세이트에 대하여 증가된 내성을 갖는 세포주 및 하이브리드를 제조하는데 친주로서 사용할 수 있다.Strains transformed according to the present invention can be used as parent strains to prepare cell lines and hybrids with increased resistance to glyphosate.

[실시예 3]Example 3

글리포세이트에 대하여 내성인, Spring colzas, Westar cultivar를 pRPA BL 410을 사용하여 불터(BOULTER) 등의 방법을 사용하여 수득한다. 이들 식물은 400g a.s/ha의 글리포세이트로 온실처리시 내성이 있었으며, 이처리로 비형질전환 식물은 파괴된다.Spring colzas, Westar cultivar, resistant to glyphosate, are obtained using a method such as BOULTER using pRPA BL 410. These plants were resistant to greenhouse treatment with 400 g a.s / ha of glyphosate, which destroyed non-transforming plants.

Claims (18)

전사방향으로, 리블로오스-1,5-디포스페이트 카르복실라제(RuBisCO) 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 전달펩티드-코딩배열, RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 N-말단 성숙 부분의 부분적 배열 및 RuBisCO유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 두번째 전달펩티드-코딩배열로 이루어진 전달펩티드-코딩 DNA 배열.In the transcription direction, the peptide-coding arrangement of the plant gene encoding the plastid-position enzyme derived from the SSU of the Ribose-1,5-diphosphate carboxylase (RuBisCO) gene, the SSU of the RuBisCO gene Peptide-coding consisting of a partial sequence of the N-terminal mature portion of the plant gene encoding the plastid-position enzyme and a second delivery peptide-coding sequence of the plant gene encoding the plastid-position enzyme derived from the SSU of the RuBisCO gene DNA array. 제1항에 있어서, 두번째 전달펩티드를 코딩하는 배열이 첫번째 전달펩티드를 코딩하는 배열이 유래된 식물과 동일한 식물로부터 유래된 전달펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein the arrangement encoding the second delivery peptide is from the same plant from which the arrangement encoding the first delivery peptide is derived. 제1항 또는 제2항에 있어서, 두번째 전달 펩티드를 코딩하는 배열이 첫번째 전달 펩티드가 유래된 식물과 다른 식물로 부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein the sequence encoding the second delivery peptide is derived from a plant different from the plant from which the first delivery peptide is derived. 제1항에 있어서, 전달 펩티드중 하나를 코딩하는 적어도 하나의 배열이 성숙 부분의 배열에 대한 원래의 배열이 유래된 유전자와 동일한 유전자로 부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein at least one of the sequences encoding one of the transfer peptides is derived from the same gene from which the original sequence for the sequence of mature portions is derived. 제1항에 있어서, 전달 펩티드중 하나를 코딩하는 적어도 하나의 배열이 옥수수 유전자로 부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein at least one array encoding one of the delivery peptides is derived from a corn gene. 제1항에 있어서, 전달 펩티드중 하나를 코딩하는 적어도 하나의 배열이 해바라기 유전자로 부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein at least one of the sequences encoding one of the delivery peptides is derived from a sunflower gene. 제1항에 있어서, 성숙 부분의 배열이 옥수수 유전자로 부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein the arrangement of the mature portions is derived from a corn gene. 제1항에 있어서, 성숙 부분의 배열이 해바라기 유전자로부터 유래된 전달 펩티드-코딩 DNA 배열.The delivery peptide-encoding DNA sequence of claim 1, wherein the arrangement of the mature portions is derived from a sunflower gene. 제8항에 있어서, 성숙 부분의 배열이 10내지 40개의 아미노산으로 이루어진 전달 펩티드-코딩 DNA 배열.10. The delivery peptide-coding DNA sequence of claim 8, wherein the mature moiety consists of 10 to 40 amino acids. 제1항에 있어서, 성숙 부분의 배열이 18 내지 33개의 아미노산으로 이루어진 전달 펩티드-코딩 DNA 배열.The delivery peptide-coding DNA sequence of claim 1, wherein the mature moiety consists of 18 to 33 amino acids. 전사방향으로, 프로모터 영역; RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 전달펩티드-코딩배열, RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 N-말단 성숙 부분의 부분적 배열 및 RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 두번째 전달펩티드-코딩배열로 이루어진 전달펩티드 영역 ; 글리포세이트 내성을 위한 코딩 배열 및 3′의 비번역 플리아데닐화 시그널 영역으로 이루어진 제초제에 대한 증가된 내성을 식물에 부여하기 위한 키메라 유전자.In the transfer direction, a promoter region; Peptide-coding arrangements of plant genes encoding plastid-position enzymes derived from SSU of RuBisCO gene, partial arrangement of N-terminal mature portions of plant genes encoding plastid-position enzymes derived from SSU of RuBisCO gene And a delivery peptide region consisting of a second delivery peptide-coding sequence of a plant gene encoding a plastid-position enzyme derived from the SSU of the RuBisCO gene; A chimeric gene for conferring increased resistance to a herbicide consisting of a coding sequence for glyphosate resistance and an untranslated pliadenylated signal region of 3 '. 제11항에 있어서, 제초제가 글리포세이트인 키메라 유전자.The chimeric gene of claim 11, wherein the herbicide is glyphosate. 제11항에 있어서, 제초제의 표적이 EPSPS인 키메라 유전자.The chimeric gene of claim 11, wherein the herbicide target is EPSPS. 제11항 내지 제13항중 어느 한 항에 있어서, 글리포세이트 내성을 코딩하는 배열이 세균 유래인 키메라 유전자.The chimeric gene according to any one of claims 11 to 13, wherein the arrangement encoding glyphosate resistance is derived from bacteria. 제11항 내지 제13항중 어느 한 항에 있어서, 글리포세이트 내성을 코딩하는 배열이 식물 유래인 키메라 유전자.The chimeric gene according to any one of claims 11 to 13, wherein the arrangement encoding glyphosate resistance is plant derived. 제11항 내지 제15항중 어느 한 항에 있어서, 키메라 유전자를 포함하는 pRPA-BL 410의 식물 형질전환용 벡터.The plant transformation vector of pRPA-BL 410 according to any one of claims 11 to 15, comprising a chimeric gene. 제16항에 따른 벡터로 형질전환된 아그로박테리움속 미생물.Agrobacterium microorganism transformed with the vector according to claim 16. 적어도 두 개의 RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 전달펩티드 영역 및 RuBisCO 유전자의 SSU로부터 유래된 플래스티드-위치 효소를 코딩하는 식물 유전자의 성숙 부분의 적어도 하나의 배열뿐 아니라 글리포세이트 내성을 코딩하는 적어도 하나의 배열 및 폴리아데닐화 시그널 영역을 각각 단리하고 내성 유전자의 전사 방향으로 조립하는 제11항 내지 제15항중 어느 한 항에 따른 키메라 유전자의 구성 방법.A delivery peptide region of a plant gene encoding a plastid-position enzyme derived from SSU of at least two RuBisCO genes and at least one arrangement of a mature portion of the plant gene encoding a plastid-position enzyme derived from SSU of RuBisCO gene The method of constructing a chimeric gene according to any one of claims 11 to 15, as well as isolating each of at least one array and polyadenylation signal region encoding glyphosate resistance and assembling in the transcriptional direction of the resistance gene.
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FR9102872A FR2673643B1 (en) 1991-03-05 1991-03-05 TRANSIT PEPTIDE FOR THE INSERTION OF A FOREIGN GENE INTO A PLANT GENE AND PLANTS TRANSFORMED USING THIS PEPTIDE.
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PT924299E (en) 2004-08-31
BR9200790A (en) 1992-11-17
IE990458A1 (en) 2000-11-15
JPH0595789A (en) 1993-04-20
DE69226466D1 (en) 1998-09-10
IL101115A (en) 1997-01-10
DE69226466T2 (en) 1999-04-01
JP3257816B2 (en) 2002-02-18
ZA921645B (en) 1992-11-25
FR2673643A1 (en) 1992-09-11
IE920690A1 (en) 1992-09-09
KR920018220A (en) 1992-10-21
EP0924299A1 (en) 1999-06-23
MX9200915A (en) 1992-09-01
EP0508909B1 (en) 1998-08-05
EP0924299B1 (en) 2004-05-19
AU652610B2 (en) 1994-09-01
DK0924299T3 (en) 2004-08-09

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